Growth and morphological response of old-forest lichens transplanted into a young and an old Picea abies forest

This experimental study focuses on why old-forest lichens such as Lobaria scrobiculata and Platismatia norvegica are scarce in younger spruce stands. Understanding the factors limiting the distribution of species is important for developing appropriate methods for forest management aiming to maintain biodiversity. A successful growth of L. scrobiculata and P. norvegica was found in the young planted stand as the rate of growth did not differ between the young stand and the old spruce forest where they naturally occurred, during 14 months of transplantation. In the young forest environment, L. scrobiculata showed a significantly higher specific thallus weight, and a slightly higher water-holding capacity. This morphological response is probably due to a higher light exposure in the young stand and consequently a higher rate of desiccation. The ubiquitous species Platismatia glauca showed a significantly higher rate of growth in the young forest than in the old forest, and a positive relationship between growth rate and light exposure was found. This transplant study has shown that the environmental conditions in younger planted forests are not necessarily unfavourable for growth of old-forest lichens. Other factors, such as limited dispersal ability and poor diaspore production, are probably important for explaining the species scarcity in younger stands.     

Defensive compound concentration in boreal lichens in response to simulated nitrogen deposition

Nitrogen (N) deposition is expected to increase in northwestern Europe the next 50–100 years. The effects of higher N availability on lichens will presumably depend on their capacity to acquire carbon (C), that is, of the timing and duration of the wet and active state. If lichens respond like plants, their C and N status may affect their concentration of carbon-based secondary compounds (CBSCs), and thus their defence against herbivores, detrimental radiation, pathogens and parasites. In the present study we have manipulated N availability and timing and duration of the metabolically active state by spraying lichen transplants in an old spruce forest with rainwater or rainwater with added N corresponding to 50 kg N ha⁻¹ yr⁻¹. The spraying was applied either at night, in the morning or at noon to also investigate the effect of timing and duration of the active state. Concentrations of N, chlorophyll a (Chl a ) and CBSCs were measured before and after one summer's spraying of 10 thalli in each of four different lichen species; Alectoria sarmentosa, Lobaria scrobiculata, Platismatia glauca , and Xanthoria aureola . The added N was readily taken up by all the lichen species. A. sarmentosa, P. glauca , and X. aureola increased their Chl a concentration in response to increased N, while L. scrobiculata increased Chl a in response to increased active time. None of the studied species reduced their concentration of secondary compounds during the experimental period, but in P. glauca the concentration of all compounds were significantly lower in N-treated thalli compared with those that got only rainwater. The results are consistent with a high degree of constitutive defence in three of four species, and we conclude that all the investigated lichens seem to have rather robust chemical defence systems despite considerable manipulation of the environmental conditions.     

Responses of the green algal foliose lichen Platismatia glauca to increased nitrogen supply

Nitrogen (N) availability and light exposure were manipulated under field conditions to study responses to altered resource supply in the green algal lichen Platismatia glauca. The lichen was fertilized with different concentrations and frequencies of ammonium, nitrate or glutamine under different light regimes for 2-3 months. Responses were followed from the intact thallus to the cellular level. Despite significant differences in overall light exposure, light conditions were not significantly different among treatments when the lichens were wet and active. Ammonium was the preferred N source, followed by glutamine and then nitrate. Thallus N concentration as well as the chlorophyll a (Chl a) concentration increased 3-4-fold at the highest ammonium concentration, while the mycobiont ergosterol concentration remained unaltered. Growth was significantly enhanced by the enhanced N supply, with the increase in dry weight varying from 3 to 30%. Variation in Chl a concentration explained 31% of this variation, suggesting a causal link to the increased growth rate. Platismatia glauca responded to increased N availability by increasing its growth rate and carbon assimilation capacity through increased investments in the photobiont cells. This suggests a tight regulation of resource investments and metabolic pathways between the symbionts of this lichen.     

Juvenile Development and Diaspore Survival in the Threatened Epiphytic Lichen Species Sticta fuliginosa, Leptogium saturninum and Menegazzia terebrata: Conclusions for in situ Conservation

Abstract: Development and growth of the three threatened epiphytic lichen species Sticta fuliginosa (Hoffm.) Ach., Leptogium saturninum (Dicks.) Nyl. and Menegazzia terebrata (Hoffm.) Massal. was investigated by low temperature scanning electron microscopy and macro‐photography. Small cotton gauze discs acting as artificial substrata were fixed with aluminium staples on the bark of selected trees and vegetative diaspores (isidia or soredia) were transferred onto these discs. The subsequent development into small thalli of up to 3 mm length was observed within the 32‐month study period. All three species produced anchoring hyphae within the first month after transplantation. Two months later 52 % of the S. fuliginosa diaspores were still on the gauze discs and after 16 months 29 % remained attached. For L. saturninum, the corresponding percentages were 46 % and 19 %, respectively. First lobes resembling adult thalli were observed after 8 to 12 months in S. fuliginosa and L. saturninum but only after 16 months in M. terebrata. All three species developed usually more than one thallus primordium (pseudomeristematic growth zone) per isidium or soredial cluster. Transplanted thallus fragments were able to fix themselves on the new substratum but in all three species large parts degenerated and fell off during the first year, particularly in S. fuliginosa. The results show that the juvenile development of the investigated species is not restricted by microclimatic factors at the study site. We therefore conclude that the juvenile development is not the restricting factor in regard to growth and population survival. Other factors, such as the competition with bryophytes, insufficient diaspore dispersal or forest management practice must account for the small population sizes. The described transplantation technique of vegetative diaspores has proved to be very useful for the augmentation of small populations without damaging the existing thalli and we suggest use of this method for in situ conservation of endangered lichen species.     

Growth and nitrogen relations in the mat-forming lichens Stereocaulon paschale and Cladonia stellaris

BACKGROUND AND AIMS: Mat-forming lichens in the genera Stereocaulon and Cladonia have ecosystem-level effects in northern boreal forests. Yet the factors affecting the productivity of mat-forming lichens are not known. The aim of the presented work was to investigate whether mat-forming lichens adapted to low N availability employ N-conserving mechanisms similar to those of vascular plants in nutrient-poor ecosystems. Specifically, the following questions were asked: (a) Do lichens translocate N from basal areas to apical growth areas? (b) Are the quantities of N translocated of ecological significance. (c) Is lichen growth dependent on tissue N concentration [N]. METHODS: Two different, but complementary, field experiments were conducted using the mat-forming N2-fixing Stereocaulon paschale and non-fixing Cladonia stellaris as model species. First, N translocation was investigated by feeding lichens with Na(15)NO3 either directly to the apex (theoretical sink) or to the basal part (theoretical source) and observing the redistribution of (15)N after a growth period. Secondly, growth and variation in [N] in thalli of different lengths was measured after a growth period. KEY RESULTS: (15)N fed to lower parts of lichen was translocated towards the growing top, but not vice versa, indicating physiologically dependent translocation that follows a sink-source relationship. In the growth experiment where thalli were cut to different lengths, the significant decrease in [N] in apices of short vs. longer thalli after a growth period is consistent with internal relocation as an ecologically important source of N. CONCLUSIONS: The presented results demonstrate that internal recycling of N occurs in both species investigated and may be ecologically important in these mat-forming lichens under field conditions. The higher nitrogen use efficiency and relative growth rate in C. stellaris in comparison with S. paschale probably enable C. stellaris to dominate the ground cover vegetation in dry boreal coniferous forests under undisturbed conditions.     

Carbon and nitrogen distribution in the green algal lichens<Emphasis Type="Italic"> Hypogymnia physodes</Emphasis> and<Emphasis Type="Italic"> Platismatia glauca</Emphasis> in relation to nutrient supply

With the aim of understanding how some lichens can survive intensive fertilization we investigated two green algal ( Trebouxia) lichens, Hypogymnia physodes (L.) Nyl. and Platismatia glauca (L.) W. Culb., and compared control (Ctr), and intensively fertilized (F) thalli. We measured total N, proteins and amino acids to assess lichen N status. Chlorophyll a indicated photosynthetic capacity and photobiont mass, ergosterol the metabolic demands of the fungus, and chitin the fungal biomass. For carbon status we measured glucose, the photobiont ( Trebouxia) export product ribitol, and the mycobiont-specific carbohydrates arabitol and mannitol. The F-thalli had 2-3 times higher protein and N concentrations, 5-10 times higher chlorophyll a concentrations, while ergosterol and chitin were doubled. The ribitol concentrations were 4-5 times higher in the F-thalli, while the fungal carbohydrates did not increase to the same extent. The amino acid arginine had increased 60-fold. The F-thalli also had a relatively higher N investment in the photobiont in relation to mycobiont tissue compared to the Ctr-thalli, probably resulting in an increased capacity for carbon assimilation, most possibly required for maintaining the higher nutrient status of the F-thalli. Arginine accumulation possibly avoided toxic effects of accumulated NH4+, albeit binding a significant fraction of assimilated carbon.     

Mollusc grazing may constrain the ecological niche of the old forest lichen Pseudocyphellaria crocata

This study reports on mollusc grazing of two epiphytic cyanobacterial lichens (Pseudocyphellaria crocata and Lobaria pulmonaria) transplanted within three Picea abies-dominated boreal rain forest stands (clear-cut, young and old forests) in west central Norway. Grazing was particularly high in transplants located in the old forest and was almost absent in clear-cut transplants. Grazing marks were absent on natural thalli on nearby spruce twigs (required creeping distance for mollusc from the ground >4 m). Transplantation of lichens from twigs to artificial transplantation frames reduced the creeping distance to 1.2 m, and caused a significant increase in grazing damage in P. crocata. Given a paired choice under transplantation, molluscs consistently preferred P. crocata and avoided L. pulmonaria, implying species-specific differences in herbivore defence. Pseudocyphellaria crocata has a much lower content of the medullary depsidones stictic and constictic acid than L. pulmonaria. Heavy grazing occurred in the P. crocata thalli lowest in these two depsidones. The upper part of the medulla hosting the photobiont was the preferred fodder for grazing molluscs. Molluscs avoided the yellow soralia in P. crocata (localised pulvinic acid), suggesting a role for pulvinic acid in preventing grazing of detached soredia and early establishment stages. The preference of P. crocata for thin spruce twigs is probably a result of a lower grazing pressure on twigs compared to e.g. deciduous stems that frequently support the better defended L. pulmonaria. Ongoing climate changes with increased annual rainfall and milder winters have presumably increased mollusc grazing, particularly in SW parts of Norway which have more species of lichen-feeding molluscs than the boreal sites studied. These temperate areas lacking natural spruce populations have recently experienced reported extinctions of the poorly defended P. crocata from rocks and deciduous stems prone to mollusc grazing. Lichen-feeding molluscs have likely played a role in these extinctions, causing spruce twigs in Atlantic boreal forests to be a last strong foothold for P. crocata in Scandinavia.     

The paradox of higher light tolerance during desiccation in rare old forest cyanolichens than in more widespread co-occurring chloro- and cephalolichens

? Desiccation tolerance was quantified in four cyanolichens (Lobaria hallii, Lobaria retigera, Lobaria scrobiculata, Pseudocyphellaria anomala), one cephalolichen (Lobaria pulmonaria) and one chlorolichen (Platismatia glauca) from xeric and mesic, open and closed North American boreal forests. ? These sympatric epiphytes were exposed to 0%, 33%, 55% and 75% relative humidity with or without medium light (200?μmol?m(-2) s(-1) ) for 7?d. Permanent and temporary photoinhibitory damage was recorded as viability measures. ? All species tolerated well the drying in darkness, but L.?hallii and L.?retigera, associated with a very humid climate, showed minor damage at the hardest drying (silica gel). Simultaneous exposure to medium light severely aggravated the drying damage at all relative humidity levels. Combined drying-light exposure was particularly devastating for the widespread chloro- and cephalolichens, whereas cyanolichens, including rare old forest species, were fairly resistant. ? The ability to recover after combined drying-light stress (this study) correlated positively with increasing species-specific water holding capacities (from the literature). Cyanolichens, depending on liquid water and large internal water storage, probably require strong drying-light resistance to handle long periods between hydration events, whereas chlorolichens can regularly maintain their photosynthetic apparatus during frequent and rapid activation by humid air on clear mornings.     

Screening the antioxidant and antimicrobial properties of the lichens Parmelia saxatilis, Platismatia glauca, Ramalina pollinaria, Ramalina polymorpha and Umbilicaria nylanderiana.

The aim of this study is to investigate in vitro antimicrobial and antioxidant activities of the methanol extracts of Parmelia saxatilis (L) Ach., Platismatia glauca (L.) W.L. Club. & C.F. Culb., Ramalina pollinaria (Wesstr.) Ach., Ramalina polymorpha (Liljeblad) Ach. and Umbilicaria nylanderiana (Zahlbr.) H. Magn. Antioxidant activity was evaluated by two separate methods: scavenging of free radical DPPH and the inhibition of linoleic acid oxidation. Extracts of Parmelia saxatilis, Platismatia glauca., Ramalina pollinaria and Ramalina polymorpha did not exert any activity in both assays, whereas those of Umbilicaria nylanderiana provided 50% inhibition at 400.2 microg/ml concentration in the former and gave 53% inhibition at 2g/l concentration. Total phenolic constituents of extracts from lichen species tested (P. saxatilis, P. glauca, R. pollinaria, R. polymorpha and U. nylanderiana) were 1.0% (w/w), 1.1% (w/w), 1.0% (w/w), 0.8% (w/w) and (3.0% w/w), respectively (as gallic acid equivalent); implying that the observed activity could be related to the amount of polar phenolics. Extracts were also found to possess antimicrobial activity against some test bacteria and fungi and yeast.     

A functional relationship between species richness of spiders and lichens in spruce

Modern forestry has created stands with even age distribution of trees and fragmentation of the habitat. In boreal forests, the effects on biodiversity within many taxa need to be examined. We tested the hypothesis that species richness of foliose and fruticose lichens and spiders is positively related in the lower canopy of spruce (Picea abies) in forests with, or without, management in central Sweden. High species richness of lichens may increase the structural complexity of the microhabitat on spruce branches, and bring a higher species richness also in the spider community. In six areas, spruce branches were sampled in old-growth and managed boreal forest stands, respectively. Forest management did not affect the species richness of spiders or lichens, but an effect due to sampling area was found in the latter taxon. There was a significant covariation between species richness of lichens and spiders, and the hypothesised positive correlation was confirmed by separate analyses for each area and combining the probabilities. Moreover, regression analysis on mean values from each site revealed a positive relationship. We conclude that species richness of lichens and spiders covary on spruce branches for functional reasons, i.e. more lichen species promotes a more diverse spider community by increasing the structural heterogeneity. Our results might provide a shortcut for assessing biodiversity in boreal forests.     

Importance of spruce swamp-forests for epiphyte diversity and flora on Picea abies in southern and middle boreal Finland

The epiphyte (lichens and bryophytes) species richness, diversity and composition on basal trunks of Picea abies in spruce swamp-forests were compared to adjacent mesic forests, on mineral soil in two southern and two middle boreal sites in old-growth forest patches in Finland The sampling was carried out along four line transects parallel to swamp-forest margin 1) in the spruce swamp-forest, 2) at the swamp-forest margin, 3) on mineral soil c 10-20 m off the swamp-forest margin and 4) on mineral soil at least 50 m off the swamp-forest margin In the two southern boreal sites there was a decreasing trend in the average species number per tree from the trees m swamp-forests (21-25 species) to the trees on mineral soil (17-18 species), whereas in the two more humid and virgin middle boreal sites a similar trend was not detected (25-28 species on all trees) There were no major differences in the epiphyte flora between the locations or study sites the dominant species occurred on nearly all sample trees The crustose lichens Cliostomum leprosum Arthoma leucopellaea and Lecanactis abietina were the most common species that were mainly confined to the swamp-forests m the southern boreal sites, C leprosum also in the middle boreal sites In addition, several rare species occurred exclusively on the swamp-forest trees Lecanactis abietina extended significantly higher on the swamp-forest trees than on the trees on mineral soil m the southern boreal sites The spruce swamp-forests proved to be one of the most important habitats for maintaining the epiphyte diversity in the boreal forest landscape     

Lichen responses to nitrogen and phosphorus additions can be explained by the different symbiont responses

? Responses to simulated nitrogen (N) deposition with or without added phosphorus (P) were investigated for three contrasting lichen species - the N-sensitive Alectoria sarmentosa, the more N-tolerant Platismatia glauca and the N(2) -fixing Lobaria pulmonaria- in a field experiment. ? To examine whether nutrient limitation differed between the photobiont and the mycobiont within the lichen, the biomass responses of the respective bionts were estimated. ? The lichenized algal cells were generally N-limited, because N-stimulated algal growth in all three species. The mycobiont was P-limited in one species (A. sarmentosa), but the growth response of the mycobionts was complex, as fungal growth is also dependent on a reliable carbon export from the photobiont, which may have been the reason for the decrease of the mycobiont with N addition in P. glauca. ? Our findings showed that P availability was an important factor when studying effects of N deposition, as P supply can both mitigate and intensify the negative effects of N on epiphytic lichens.     

CO2 stimulation and response mechanisms vary with light supply in boreal conifers

北方针叶树CO₂ 的刺激和响应机制随光强而变化 黑云杉(Picea mariana [Mill.] B.S.P.)和白云杉(Picea glauca [Moench] Voss.)是同属物种,两者都是适度耐阴,并且在北美北方针叶林中广泛分布。为了了解光照对CO₂ 浓度升高的生理生态反应的影响,在三种光照条件下(温室中光照设置为100%、50%和30%)将一年生的两种幼苗暴露在360和720 µmol mol^–1 浓度的CO₂环境中,测定了其中后期叶面气体交换量。研究结果表明,CO₂的浓度升高提高了净光合速率 (P_n)和光合水分利用效率,但降低了气孔导度和蒸腾作用。CO₂对光合作用的刺激在50%光照下最大, 在100%光照下最小。光合作用、最大羧化速率(V_cmax)和光饱和电子传递速率(J_max)均随光照强度的 降低而降低。升高的CO₂在所有光照处理中显著降低了V_cmax,在生长季节中期,两种云杉的V_cmax均显著 降低,但在生长季节后期,当光照达到30%时,这一影响变得不明显,而且黑云杉的响应大于白云杉。CO₂ 浓度升高也降低了白云杉的J_max,但在生长季后期30%光照时,这种影响变得不显著。但CO₂ 浓度升高对 黑云杉的影响随时间而变化。在所有光照处理中,CO₂ 浓度升高降低了黑云杉生长中期的J_max,且在生长后 期30%光照时影响不显著,但在100%和50%光照时,J_max升高。这些研究结果表明,两个树种植物都受益于CO₂ 浓度的升高,但它们的响应机制随着光照的增加而变化：即在100%和50%光照下,它们的响应主要是生理上的,而在30%光照下,它们的响应主要是形态上的。     

Recovery of pristine boreal forest floor community after selective removal of understorey, ground and humus layers

In boreal spruce forests that rarely experience extensive disturbances, fine-scale vegetation gaps are important for succession dynamics and species diversity. We examined the community implications of fine-scale gap disturbances by selective removal of vegetation layers in a pristine boreal spruce forest in Northern Finland. The aim was to investigate how the speed of recovery depends on the type of disturbance and the species growth form. We also wanted to know if there appeared changes in species composition after disturbance. Five different treatments were applied in the study: Control, removal of the ground layer (bryophytes and lichens), removal of the understorey layer (dwarf shrubs, herbs and graminoids), removal of both the ground and understorey layers, and complete removal of the vegetation and humus layers above the mineral soil. The vegetation recovery was monitored in terms of cover and species numbers over a 5-year period. Understorey layer cover, composed mainly of clonal dwarf shrubs, recovered completely in 4 years in treatments where the humus layer remained intact, whereas ground layer cover did not reach the control level in plots from where bryophytes and lichens were removed. Recovery was faster in terms of species number than species cover. Bryophytes, graminoids and dominant dwarf shrubs appeared in all disturbed plots quickly after disturbance. Seedlings of trees appeared exclusively in disturbed plots. Graminoids dominated after the removal of humus layer. The results indicate that the regeneration of forest floor after small gap disturbance occurs mainly by re-establishment of the dominant species. Although destruction of the humus layer leaves a long-lasting scar to the forest floor, exposing of mineral soil may enhance the sexual reproduction of dominant species and the colonization of weaker competitors.     

Effects of enhanced UV-B radiation in the field on the concentration of phenolics and chlorophyll fluorescence in two boreal and arctic–alpine lichens

Lichens constitute a prominent part of the vegetation at high latitudes and altitudes, but the effects of UV-B radiation on these symbiotic organisms are not well known. In a northern boreal site (Abisko, northern Sweden), the usnic acid-producing lichens Flavocetraria nivalis and Nephroma arcticum were exposed to enhanced UV-B radiation, corresponding to 25% ozone depletion, for two and one growing seasons, respectively. They were compared with lichens grown under ambient UV-B and harvested fresh from the field. The treated thalli of F. nivalis had been transplanted from a site 24 km from the treatment site. From this source locality, untreated thalli were also harvested. Enhanced UV-B did not affect concentrations of usnic acid and the two depsides phenarctin and nephroarctin. A gradual decline of usnic acid, probably coupled to unusually long periods of dry, sunny weather, was observed both under enhanced and ambient UV-B and in untreated thalli. Photosystem II efficiency in both species was slightly reduced by enhanced UV-B. However, differences between seasons were larger than differences between treatments, which indicate that UV-B effects are minor in comparison to other climatic variables. Concentrations of UV-B-absorbing phenolics in lichens do not show a simple relationship to UV-B dose and therefore cannot be used as bioindicators of UV-B levels.     

Effects of Retention Felling on Epixylic Species in Boreal Spruce Forests in Southern Finland

We studied the effects of patch retention felling and soil scarification by harrowing on the coverage and species richness of epixylic species in boreal Norway spruce (Picea abies) forests in Southern Finland. The epixylics were investigated from both the retention tree groups (RTGs) and the surrounding felling areas before and after fellings and after scarification on consecutive years. The cover percentage of all included species groups was shown to decrease after the felling, especially in the felling areas (vascular plants, ?0.4%; mosses, ?27.8%; liverworts, ?4.0%; and lichens,?2.1%). The decrease was considerable also in the RTGs. The amount of dead moss increased in both the areas indicating microclimatic changes. Species richness also declined rapidly after the first year, especially in the felling areas (vascular plants, ?2.2%; mosses, ?27.3%; liverworts, ?30.3%; and lichens, ?22.9%). Scarification also decreases covers and species richness of bryophytes. After the second year, the covers of the species groups generally started to regain, especially in the untreated RTGs. The size of RTG was in positive correlation with the total species number. Another main result indicates that it is possible to maintain much higher initial vegetation abundance and diversity in the RTGs than in the felling areas. Coarse woody debris formed by the frequent tree uprootings may also enhance the long‐term survival of epixylics over forest regeneration period. RTGs should be at least 10 times larger than the size used in current Finnish forestry, so that they could function as species refugia.     

Seasonal partitioning of growth into biomass and area expansion in a cephalolichen and a cyanolichen of the old forest genus Lobaria

Growth in two old forest lichens was studied to evaluate how temporal (seasonal) and spatial (aspect-wise) partitioning of biomass and area growth respond to seasonal changes in light and climate. We monitored relative growth rates during annual courses in the cephalolichen Lobaria pulmonaria and the cyanolichen Lobaria scrobiculata transplanted in boreal clear-cut to five fixed aspects in winter, spring, summer, and autumn. For each annual set, growth was quantified in January-March, April-June, July-September and October-December. Mean biomass and area increased in all seasons, but growth was highest in July-September. Mass growth did not follow area increment during a year. As a result, mass per area (specific thallus mass (STM)) declined (L. scrobiculata) or stayed constant (L. pulmonaria) in the dark, humid October-December season, whereas it strongly increased in the dry, sunny April-June season. Aspect influenced growth in species-specific ways. Seasonality in biomass growth mainly followed light availability, whereas area growth was strongest during humid seasons. The substantial STM changes across seasons, species, and aspects can be explained as passive responses to seasonal climate. However, as STM, according to the literature, is a driver of water storage, recorded changes probably improve fitness by prolonging hydration in places or during times with high evaporative demands.     

Growth and vitality of epiphytic lichens

 We tested the hypothesis that changed microclimate at induced forest edges causes reduced growth of epiphytic lichens. Two foliose, green algal lichens were transplanted to the lower canopy of a mature Picea abies forest at six distances (2, 6.25, 12.5, 25, 50 and 100 m) from a clearcut. The biomass growth in Platismatia glauca (6.2% in 16 months) was 41% higher than in Lobaria pulmonaria (4.4%). We found no growth reduction near the forest edge. In contrast, the highest growth in both species occurred within 12 m from the edge. Further, fluorescence and chlorophyll measurements showed that lichen vitality was unaffected by distance from edge. The light intensity was 4.3 times higher at the edge than in the interior during the growing season, but there were only minor differences in air temperature and relative humidity. Monitoring of thallus water content revealed clear differences in both number and length of wetting and drying cycles. However, the total time with water content sufficient for photosynthetic activity was only slightly higher at the edge. The data thus indicate that our gradient in microclimate was too small to significantly affect lichen growth, and that lichens are largely metabolically inactive when large edge-interior contrasts in microclimate occur. Lichen response to forest edge microclimate results from intricate interactions among several biotic and abiotic factors. Linking data on lichen growth, microclimate and thallus water content with physiological measurements provides a framework for future studies of the mechanisms behind abiotic edge effects. Received: 15 April 1996 / Accepted: 21 June 1996     

Cytotoxic activity of some lichen extracts on murine and human cancer cell lines

Eight lichens were extracted successively with n-hexane, diethyl ether and methanol using a Soxhlet process. The cytotoxic activity of the 24 lichen extracts was evaluated in vitro using two murine (the L1210: lymphocytic leukaemia, and the 3LL: Lewis lung carcinoma) and four human (the K-562: chronic myelogenous leukaemia, the U251: glioblastoma, the DU145: prostate carcinoma, and the MCF7: breast adenocarcinoma) cancer cell lines and non-cancerous cells, the Vero cell line (African green monkey kidney cell line). The MTT assay revealed significant cytotoxicity (IC50 < or = 20 microg/ml) on one of the tested cancer cell lines for at least one extract of each lichen species. Some extracts of Cladonia convoluta, Cladonia rangiformis, Parmelia caperata, Platismatia glauca and Ramalina cuspidata demonstrated interesting activities particularly on human cancer cell lines as good selectivity indices were recorded (SI > 3).     

Growth of foliose lichens: a review

This review considers various aspects of the growth of foliose lichens including early growth and development, variation in radial growth rate (RaGR) of different species, growth to maturity, lobe growth variation, senescence and fragmentation, growth models, the influence of environmental variables, and the maintenance of thallus symmetry. The data suggest that a foliose lichen thallus is essentially a ‘colony’ in which the individual lobes exhibit a considerable degree of autonomy in their growth processes. During development, recognisable juvenile thalli are usually formed by 15 months to 4 years while most mature thalli exhibit RaGR between 1 and 5 mm yr⁻¹. RaGR within a species is highly variable. The growth rate-size curve of a foliose lichen thallus may result from growth processes that take place at the tips of individual lobes together with size-related changes in the intensity of competition for space between the marginal lobes. Radial growth and growth in mass is influenced by climatic and microclimatic factors and also by substratum factors such as rock and bark texture, chemistry, and nutrient enrichment. Possible future research topics include: (1) measuring fast growing foliose species through life, (2) the three dimensional changes that occur during lobe growth, (3) the cellular changes that occur during regeneration, growth, and division of lobes, and (4) the distribution and allocation of the major lichen carbohydrates within lobes.